Henrike Hultsch

Temporal performance roles during vocal interactions in nightingales (Luscinia megarhynchos B.)

SummaryThe temporal patterning of vocal interactions between territorial nightingales (Luscinia megarhynchos B). was investigated during nocturnal dyadic singing. We distinguished three forms of temporal performance roles (Table 1; Fig. 3): inserter (preferred song start 0.5–1.0 s after offset of a neighbours song), overlapper (preferred song start 0.5–1.0 s after onset of a neighbours song), and Autonomous bird (no evidence for temporal responses adjusted to the timing of a neighbours singing; Fig. 4). With the beginning of the daily dawn chorus, mutual temporal adjustment of song performances could no longer be ascertained (Fig. 2). To test the flexibility of song onset timing, we presented subjects with three playback programs (I, II, III) simulating a non-flexible conspecific bird. Sequences of songs recorded from a stranger that shared no song types with the subjects were used; intersong pauses were unaltered in I, lengtened in II, and shortened in III. Results confirmed that the timing of song onset can be influenced by the temporal patterning of auditory stimuli (Table 1; Fig. 5): Inserters and overlappers sang at a slower rate during program II than during program I. During program III, which by accelerated stimulus succession caused frequent overlap of songs, inserters and overlappers interrupted their singing behaviour (Fig. 6). This suggests that overlapping, which results in signal interference, is of some functional significance in vocal interactions.

Repertoire sharing and song-post distance in nightingales (Luscinia megarhynchos B.)

SummaryThe relationship was studied between song-post distances and the extent of vocal repertoire sharing in 34 territorial nightingales settling in six homogeneously structured habitats. Repertoires were compared on the basis of shared song types and distances were measured between nocturnal song posts of first-order and higher-order neighbours. Our results showed that male nightingales shared fewer song types with very close and more distant neighbours than with neighbours at intermediate distances. This distribution is explained by the interaction betwee repelling and attracting components, which depends on the distance between song posts.

Song repertoire size is correlated with body measures and arrival date in common nightingales, Luscinia megarhynchos

In songbirds, the individual features of a males song play a role in female choice and male–male interactions. Therefore, it has been hypothesized that the song characteristics of an individual should be correlated with its quality and this has been shown for some species. The species studied so far have only small or medium-sized song repertoires, leaving open the question of whether, in species with large repertoires, song characteristics such as repertoire size are related to male quality. We investigated the correlations between body measures, song characteristics and arrival dates of common nightingales, a songbird species in which males have extraordinarily large song type repertoires. We found that birds with large repertoires had longer wings and were heavier, and arrived earlier, than those with small repertoires. Other song characteristics, such as song diversity and proportion of whistle songs, were not significantly correlated with body measures. When we correlated body measures with arrival date, we found that birds that arrived earlier were heavier and tended to be in better condition. These findings provide supporting evidence that repertoire size is an indicator of a males quality.

Chapter 3 – Learning to sing

The development of singing in birds ranks as a prime biological model for understanding how nature and nurture interact, and how they jointly affect the growth of behavioral competence. Research on this subject benefits from contributions from many different disciplines, including neurobiology and genetics, as well as behavioral biology. There are further advantages. Behavioral studies offer the prospect of uncovering aspects of song development at three different levels. Besides an investigation of so-called developmental trajectories, including rules that describe the many transformations of a juveniles vocal utterances, there are two kinds of reference patterns to which each vocalization can be compared; the stimulus patterns that a bird has experienced earlier, and the target songs that he finally performs as an adult. Learning to sing takes some time. Usually it begins when a young bird leaves the nest, and then continues for several months. It comes to an end only shortly before the bird establishes his own territory and flags its turf by singing. Although some species also learn new songs later in life, the first year of song development is crucial in all songbirds. Their many accomplishments can be subdivided into two major stages: the auditory phase of song memorization, and the motor phase of song development. The chapter treats the various aspects of learning to sing in some detail, keeping as a framework of typical biological concepts that are often used to explain the behavioral accomplishments of organisms, either as a consequence of evolutionary adaptation, or as mechanisms to solve a particular problem successfully.

Early experience can modify singing styles: evidence from experiments with nightingales, Luscinia megarhynchos

Abstract The way in which song repertoires are performed, though modifiable by degree, has been considered to be a species-typical trait. This study examines whether it can be influenced by learning. During their early sensitive phase of song acquisition, hand-raised nightingales were tutored with strings of master song-types in which song-types were arranged in different bout structures. In one design (control), strings with the species-typical song-type sequencing were presented (i.e. versatile bout structure, no song-type repetitions: A,B,C…). In the other design song-types were arranged in a bout structure atypical for the species (i.e. song-type repetitions: D,D,D,… E,E,E…). Subjects were exposed either to both designs (experimental group) or to the versatile design only (control group). The crystallized singing of experimental subjects at 11 months revealed that they had acquired serial information about the tutored strings. Song-types acquired from the repetitive tutoring design were repeated significantly more often than those acquired from the versatile design. Furthermore, the bout length of song-type renditions showed a relationship to the bout length of the respective master song-types upon presentation. The results provide evidence for an acquisition of repertoire performance programmes through learning, which may modulate species-typical singing styles.

Long-term persistence of song performance rules in nightingales ( Luscinia megarhynchos ): a longitudinal field study on repertoire size and composition

Common nightingales (Luscinia megarhynchos) are among those bird species that possess an extremely large repertoire and perform it in a versatile singing style. Thereby, repertoire size, composition, and performance differs considerably among individuals. In this longitudinal field study, we investigated the long-term stability of these differences in the song characteristics of free-ranging nightingales. We determined the repertoire characteristics for nine adult male individuals in two successive years (three of these individuals were investigated over the course of three years) and compared these to similar measurements obtained from comparisons of song samples of different birds. Comparisons revealed remarkable differences among males, but we did not find systematic differences in the song performance of birds in successive years. Instead, song characteristics were remarkably stable within successive years. The long-term persistence of individual song characteristics suggests that they are not related to dynamically changing individual attributes, but may reflect long-term storage of information during song acquisition as juveniles. In addition, we found that the repertoire performance of adult nightingales allows fine-tuned vocal interactions among several neighbouring males.

How songbirds deal with large amounts of serial information: retrieval rules suggest a hierarchical song memory

Abstract. Many songbirds develop remarkably large vocal repertoires, and this has prompted questions about how birds are able to successfully learn and use the often enormous amounts of information encoded in their various signal patterns. We have studied these questions in nightingales (Luscinia megarhynchos), a species that performs more than 200 different types of songs (strophen), or more than 1000 phonetically different elements composing the songs. In particular, we investigated whether and how both song repertoires and song performance rules of nightingales were coded by auditory stimuli presented in serial learning experiments. Evaluation of singing episodes produced by our trained birds revealed that nightingales cope well with an exposure to even long strings of master song-types. They can readily acquire information encoded within and between the different master songs, and they memorize, for example, which master song-types they have experienced in the same learning context. Imitations of such song-types form distinct sequential associations that are termed “context groups”. Additionally, nightingales develop other song-type associations that are smaller in size and termed “package groups”. Package formation results from constraints of the acquisition mechanisms which obviously lead to a segmentation of auditorily perceived master song sequences. Further experimentation validated that the song memory of nightingales is organized in a hierarchical manner and holding information about “context groups” composed of packages, “package groups” composed of songs, and songs composed of song elements. The evidence suggests that implementation of such a hierarchical organization facilitates a quick retrieval of particular songs, and thereby provides an essential prerequisite for a functionally appropriate use of large vocal repertoire is in songbirds.

Latent song type memories are accessible through auditory stimulation in a hand-reared songbird

The production of learned vocalizations such as in birdsong is often used to judge whether stimuli had been memorized upon their presentation. However, failures in the imitation of certain song patterns may also reflect impaired development of motor programmes or impaired memory retrieval rather than failures in stimulus memorization during auditory acquisition. To study this issue, we confronted adult hand-reared nightingales, Luscinia megarhynchos, with interactive playback experiments and used vocal matching as a behavioural tool to investigate their song type memories. Vocal matching is a common pattern-specific response that songbirds use in territorial countersinging. We distinguished two forms of pattern-specific matching: (1) song type matching (i.e. a bird replied with the same song type as the stimulus song), and (2) song group matching (i.e. the bird replied with a different song type which was, however, sequentially associated with the playback song presented earlier, i.e. during the tutoring). Some subjects used both song type and song group matching in response to song types they had not imitated from the tutor programme prior to the playback experiments. Our results indicate that nightingales store more song types in their sensory phase than they spontaneously recall from memory as adults. That is, memories of song types that were not performed in overt behaviour could be activated by vocal interactions, here induced by the interactive playback.

Responses to playback of whistle songs and normal songs in male nightingales: effects of song category, whistle pitch, and distance

Abstract. In most species of song birds, males develop song repertoires of several different songs. Among this variety, different songs may be used differently in communication and, thus, may have different functions. Here we studied vocal responses to playback of structurally different songs in male territorial nightingales (Luscinia megarhynchos). Nightingales have enormous vocal repertoires of about 200 song types. Among these songs, one category, whistle songs, sticks out syntactically and acoustically. Here we tested whether or not male nightingales match whistle songs with whistle songs and, if so, whether they also match the pitch of the broadcast whistles. Furthermore, we also tested if nightingales treat whistle songs as a separate category of songs. We conducted interactive playback experiments on nocturnal song in which each male received three playback treatments that differed in the number of whistle songs broadcast. Males responded differently during playbacks by singing significantly more whistle songs when the playback tapes contained many whistle songs than when they contained no whistle songs. Males also frequently matched the pitch of the broadcast whistle songs. In contrast to responses during playback, after the playback terminated males sang more whistle songs when no whistle songs were broadcast than when many whistle songs were broadcast. These findings suggest that whistle songs have a specific signal value and that nightingales treat them as a special song category. Electronic supplementary material to this paper can be obtained by using the Springer Link server located at http://dx.doi.org/10.1007/s00265-002-0511-1.

The use of network analysis to study complex animal communication systems: a study on nightingale song

The singing of song birds can form complex signal systems comprised of numerous subunits sung with distinct combinatorial properties that have been described as syntax-like. This complexity has inspired inquiries into similarities of bird song to human language; but the quantitative analysis and description of song sequences is a challenging task. In this study, we analysed song sequences of common nightingales (Luscinia megarhynchos) by means of a network analysis. We translated long nocturnal song sequences into networks of song types with song transitions as connectors. As network measures, we calculated shortest path length and transitivity and identified the ‘small-world’ character of nightingale song networks. Besides comparing network measures with conventional measures of song complexity, we also found a correlation between network measures and age of birds. Furthermore, we determined the numbers of in-coming and out-going edges of each song type, characterizing transition patterns. These transition patterns were shared across males for certain song types. Playbacks with different transition patterns provided first evidence that these patterns are responded to differently and thus play a role in singing interactions. We discuss potential functions of the network properties of song sequences in the framework of vocal leadership. Network approaches provide biologically meaningful parameters to describe the song structure of species with extremely large repertoires and complex rules of song retrieval.